Rodents change acorn dispersal behaviour in response to ungulate presence

Small rodents are prominent seed predators, but they also favour plant recruitment as seed dispersers. The direct interactions of ungulates on plants are more one‐sided and negative, as they mainly reduce plant recruitment through predation on seeds and seedlings. The effects of small rodents and ungulates on plant recruitment have been considered and studied as independent episodes within plant regeneration cycles. However, ungulate–rodent interactions and their potential effects on plant regeneration have not been considered so far. A number of studies have recently documented ungulate effects on the abundance, diversity and spatial distribution of small rodents. Here, we hypothesize that ungulates may also affect rodent seed dispersal behaviour. We monitored acorn dispersal by small rodents (Mus spretus and Apodemus sylvaticus) in oak woodlands with and without exclosures for large ungulates, mainly red deer, Cervus elaphus, and wild boar, Sus scrofa. The study was carried out in a typical Mediterranean Holm oak, Quercus ilex, forest throughout the acorn fall season in 2003 and 2004. We found that, in both years, the proportion of acorns cached and not recovered in the short‐term was, on average, lower in the presence (1.4%) than in the absence (19.9%) of ungulates. Acorn dispersal distances were not affected by ungulate presence in either year. However, ungulates had an effect on the spatial distribution of dispersed seeds; rodents apparently avoided shrubs as caching sites in both years. This result was interpreted as a behavioural response to reduce the risk of cache pilferage by conspecifics, which are closely associated with shrubs in presence, but not in absence, of ungulates. Potential effects of different densities of rodents or predators were discarded, as none of them differed between the areas with and without ungulates. The present study found significant interactions between heterospecific seed and seedling consumers that had been considered as independent episodes within tree regeneration cycles. As a result of such interactions, ungulates may have negative indirect effects on oak recruitment by reducing (1) acorn caching frequency, and (2) the proportion of acorns cached under shrubs, key nurse‐plants for the establishment of Holm oak seedlings in Mediterranean areas.     

Ungulates increase forest plant species richness to the benefit of non‐forest specialists

Large wild ungulates are a major biotic factor shaping plant communities. They influence species abundance and occurrence directly by herbivory and plant dispersal, or indirectly by modifying plant‐plant interactions and through soil disturbance. In forest ecosystems, researchers’ attention has been mainly focused on deer overabundance. Far less is known about the effects on understory plant dynamics and diversity of wild ungulates where their abundance is maintained at lower levels to mitigate impacts on tree regeneration. We used vegetation data collected over 10 years on 82 pairs of exclosure (excluding ungulates) and control plots located in a nation‐wide forest monitoring network (Renecofor). We report the effects of ungulate exclusion on (i) plant species richness and ecological characteristics, (ii) and cover percentage of herbaceous and shrub layers. We also analyzed the response of these variables along gradients of ungulate abundance, based on hunting statistics, for wild boar (Sus scrofa), red deer (Cervus elaphus) and roe deer (Capreolus capreolus). Outside the exclosures, forest ungulates maintained higher species richness in the herbaceous layer (+15%), while the shrub layer was 17% less rich, and the plant communities became more light‐demanding. Inside the exclosures, shrub cover increased, often to the benefit of bramble (Rubus fruticosus agg.). Ungulates tend to favour ruderal, hemerobic, epizoochorous and non‐forest species. Among plots, the magnitude of vegetation changes was proportional to deer abundance. We conclude that ungulates, through the control of the shrub layer, indirectly increase herbaceous plant species richness by increasing light reaching the ground. However, this increase is detrimental to the peculiarity of forest plant communities and contributes to a landscape‐level biotic homogenization. Even at population density levels considered to be harmless for overall plant species richness, ungulates remain a conservation issue for plant community composition.     

Antelope mating strategies facilitate invasion of grasslands by a woody weed

Intra and interspecific variation in frugivore behaviour can have important consequences for seed dispersal outcomes. However, most information comes from among‐species comparisons, and within‐species variation is relatively poorly understood. We examined how large intraspecific differences in the behaviour of a native disperser, blackbuck antelope Antilope cervicapra, influence dispersal of a woody invasive, Prosopis juliflora, in a grassland ecosystem. Blackbuck disperse P. juliflora seeds through their dung. In lekking blackbuck populations, males defend clustered or dispersed mating territories. Territorial male movement is restricted, and within their territories males defecate on dung‐piles. In contrast, mixed‐sex herds range over large areas and do not create dung‐piles. We expected territorial males to shape seed dispersal patterns, and seed deposition and seedling recruitment to be spatially localized. Territorial males had a disproportionately large influence on seed dispersal. Adult males removed twice as much fruit as females, and seed arrival was disproportionately high on territories. Also, because lek‐territories are clustered, seed arrival was spatially highly concentrated. Seedling recruitment was also substantially higher on territories compared with random sites, indicating that the local concentration of seeds created by territorial males continued into high local recruitment of seedlings. Territorial male behaviour may, thus, result in a distinct spatial pattern of invasion of grasslands by the woody P. juliflora. An ex situ experiment showed no beneficial effect of dung and a negative effect of light on seed germination. We conclude that large intraspecific behavioural differences within frugivore populations can result in significant variation in their effectiveness as seed dispersers. Mating strategies in a disperser could shape seed dispersal, seedling recruitment and potentially plant distribution patterns. These mating strategies may aid in the spread of invasives, such as P. juliflora, which could, in turn, negatively influence the behaviour and ecology of native dispersers.     

Density dependence of daily activity in three ungulate species

Daily activity in herbivores reflects a balance between finding food and safety. The safety‐in‐numbers theory predicts that living in higher population densities increases safety, which should affect this balance. High‐density populations are thus expected to show a more even distribution of activity—that is, spread—and higher activity levels across the day. We tested these predictions for three ungulate species; red deer (Cervus elaphus), roe deer (Capreolus capreolus), and wild boar (Sus scrofa). We used camera traps to measure the level and spread of activity across ten forest sites at the Veluwe, the Netherlands, that widely range in ungulate density. Food availability and hunting levels were included as covariates. Daily activity was more evenly distributed when population density was higher for all three species. Both deer species showed relatively more feeding activity in broad daylight and wild boar during dusk. Activity level increased with population density only for wild boar. Food availability and hunting showed no correlation with activity patterns. These findings indicate that ungulate activity is to some degree density dependent. However, while these patterns might result from larger populations feeling safer as the safety‐in‐numbers theory states, we cannot rule out that they are the outcome of greater intraspecific competition for food, forcing animals to forage during suboptimal times of the day. Overall, this study demonstrates that wild ungulates adjust their activity spread and level based on their population size.     

What Cues Do Ungulates Use to Assess Predation Risk in Dense Temperate Forests?

Anti-predator responses by ungulates can be based on habitat features or on the near-imminent threat of predators. In dense forest, cues that ungulates use to assess predation risk likely differ from half-open landscapes, as scent relative to sight is predicted to be more important. We studied, in the Białowieża Primeval Forest (Poland), whether perceived predation risk in red deer (Cervus elaphus) and wild boar (Sus scrofa) is related to habitat visibility or olfactory cues of a predator. We used camera traps in two different set-ups to record undisturbed ungulate behavior and fresh wolf (Canis lupus) scats as olfactory cue. Habitat visibility at fixed locations in deciduous old growth forest affected neither vigilance levels nor visitation rate and cumulative visitation time of both ungulate species. However, red deer showed a more than two-fold increase of vigilance level from 22% of the time present on control plots to 46% on experimental plots containing one wolf scat. Higher vigilance came at the expense of time spent foraging, which decreased from 32% to 12% while exposed to the wolf scat. These behavioral changes were most pronounced during the first week of the experiment but continuous monitoring of the plots suggested that they might last for several weeks. Wild boar did not show behavioral responses indicating higher perceived predation risk. Visitation rate and cumulative visitation time were not affected by the presence of a wolf scat in both ungulate species. The current study showed that perceived predation risk in red deer and wild boar is not related to habitat visibility in a dense forest ecosystem. However, olfactory cues of wolves affected foraging behavior of their preferred prey species red deer. We showed that odor of wolves in an ecologically equivalent dose is sufficient to create fine-scale risk factors for red deer.     

Inter-specific synchrony of two contrasting ungulates: wild boar (Sus scrofa) and roe deer (Capreolus capreolus)

Very few studies on ungulates address issues of inter-specific synchrony in population responses to environmental variation such as climate. Depending on whether annual variation in performance of ungulate populations is driven by direct or indirect (trophic) interactions, very different predictions regarding the pattern of inter-specific synchrony can be derived. We compared annual autumn body mass variation in roe deer (Capreolus capreolus) and wild boar (Sus scrofa) from Poland over the period 1982–2002, and related this to variation in winter and summer climate and plant phenological development [the Normalized Difference Vegetation Index (NDVI), derived from satellites]. Roe deer fawns (∼1.3 kg increase from year 1982 to 2002) and yearlings both increased markedly in mass over years. There was also an increase for wild boar mass over years (∼4.2 kg increase for piglets from 1982 to 2002). Despite our failure to link annual body mass to spring or winter conditions or the NDVI, the body mass of roe deer and wild boar fluctuated in synchrony. As this was a field roe deer population, and since wild boar is an omnivore, we suggest this may be linked to annual variation and trends in crop structure (mainly rye). We urge future studies to take advantage of studying multiple species in order to gain further insight into processes of how climate affect ungulate populations. Electronic supplementary material Supplementary material is available in the online version of this article at and is accessible for authorized users.     

Mammalian seed dispersal in Cantabrian woodland pastures: Network structure and response to forest loss

We evaluated the role of wild large mammals as dispersers of fleshy-fruited woody plants in woodland pastures of the Cantabrian range (N Spain). By searching for seeds in mammal scats across four localities, we addressed how extensive seed dispersal was in relation to the fleshy-fruited plant community, and applied a network approach to identify the relative role of mammal species in the seed dispersal process. We also tested the response of mammalian dispersers to forest availability at increasing spatial scales. Five carnivores and three ungulates dispersed seeds of eight fleshy-fruited trees and shrubs. Mammalian seed dispersal did not mirror community-wide fruit availability, as abundant fruiting trees were scarce whereas thorny shrubs were over-represented among dispersed species. The dispersal network was dominated by bramble (Rubus ulmifolius/fruticosus), the remaining plants being rarer and showing more restricted disperser coteries. Fox (Vulpes vulpes), badger (Meles meles), and wild boar (Sus scrofa) dispersed mostly bramble, whereas martens (Martes sp.) dispersed mostly wild rose (Rosa sp.). Ungulates occasionally dispersed holly (Ilex aquifolium) and hawthorn (Crataegus monogyna). The empirical network reflected a skewed distribution of interactions and some functional complementarity (as judged from the low levels of connectance and nestedness), but also some degree of specialization. Mammals overused uncovered microsites for seed deposition, and increased their disperser activity in those landscape sectors devoid of forest. Combined with previous findings on avian seed dispersal, this study suggest a strong functional complementarity coming from the low overlap in the main plant types that mammals and birds disperse – thorny shrubs and trees, respectively – and the differential patterns of seed deposition, with mammals mostly dispersing into deforested areas, and birds into forest-rich landscapes.     

Endozoochorous seed dispersal by free-ranging herbivores in an abandoned landscape

Seed dispersal is a crucial process for the dynamics and maintenance of plant populations. Free-ranging animals are effective dispersal vectors because they can move between similar habitats and transport seeds into favourable environments. Dung samples from two species of common free-ranging mammals—deer and wild boar—were used to study endozoochorous dispersal of seeds in a military training area in western Bohemia. The area was abandoned after WWII, and the military training area was established in 1953. The vegetation consists of shrublands and dry grasslands. Data on the local species pool of grassland herbs and forbs were collected to compare the characteristics of dispersed versus non-dispersed plants. Deer and wild boar dispersed 84 plant species; however, species composition of seedlings emerging from dung samples showed significant differences between dispersal vectors and notable change across the growing season. 80% of all seedlings extracted from the dung samples belonged to stinging nettle, Urtica dioica. From trait analyses, seeds of endozoochorous plants had a higher longevity index in the soil seed bank than non-endozoochorous plants and more often possessed a mucilaginous surface. Our results show that deer and boar are successful, though not substitutable dispersers.     

Disentangling herbivore impacts on Populus tremuloides: a comparison of native ungulates and cattle in Canada’s Aspen Parkland

Ungulates impact woody species’ growth and abundance but little is understood about the comparative impacts of different ungulate species on forest expansion in savanna environments. Replacement of native herbivore guilds with livestock [i.e., beef cattle (Bos taurus)] has been hypothesized as a factor facilitating trembling aspen (Populus tremuloides Michx.) encroachment into grasslands of the Northern Great Plains. We used a controlled herbivory study in the Parklands of western Canada to compare the impact of native ungulates and cattle on aspen saplings. Native ungulate treatments included a mixed species guild and sequences of herbivory by different ungulates [bison (Bison bison subsp. bison), elk (Cervus elaphus) then deer (Odocoileus hemionus); or deer, elk, then bison]. Herbivory treatments were replicated in three pastures, within which sets of 40 marked aspen saplings (<1.8 m) were tracked along permanent transects at 2-week intervals, and compared to a non-grazed aspen stand. Stems were assessed for mortality and incremental damage (herbivory, leader breakage, stem abrasion and trampling). Final mortality was greater with exposure to any type of herbivore, but remained similar between ungulate treatments. However, among all treatments, the growth of aspen was highest with exposure only to cattle. Herbivory of aspen was attributed primarily to elk within the native ungulate treatments, with other forms of physical damage, and ultimately sapling mortality, associated with exposure to bison. Overall, these results indicate that native ungulates, specifically elk and bison, have more negative impacts on aspen saplings and provide evidence that native and domestic ungulates can have different functional effects on woody plant dynamics in savanna ecosystems.     

Temporal dynamics of seed excretion by wild ungulates: implications for plant dispersal

Dispersal is a key process in metapopulation dynamics as it conditions species' spatial responses to gradients of abiotic and biotic conditions and triggers individual and gene flows. In the numerous plants that are dispersed through seed consumption by herbivores (endozoochory), the distance and effectiveness of dispersal is determined by the combined effects of seed retention time in the vector's digestive system, the spatial extent of its movements, and the ability of the seeds to germinate once released. Estimating these three parameters from experimental data is therefore crucial to calibrate mechanistic metacommunity models of plant–herbivore interactions. In this study, we jointly estimated the retention time and germination probability of six herbaceous plants transported by roe deer (Capreolus capreolus), red deer (Cervus elaphus), and wild boar (Sus scrofa) through feeding experiments and a Bayesian dynamic model. Retention time was longer in the nonruminant wild boar (>36 h) than in the two ruminant species (roe deer: 18–36 h, red deer: 3–36 h). In the two ruminants, but not in wild boar, small and round seeds were excreted faster than large ones. Low germination probabilities of the excreted seeds reflected the high cost imposed by endozoochory on plant survival. Trait‐mediated variations in retention time and germination probability among animal and plant species may impact plant dispersal distances and interact with biotic and abiotic conditions at the release site to shape the spatial patterns of dispersed plant species.     

Environmental factors shaping ungulate abundances in Poland

Population densities of large herbivores are determined by the diverse effects of density-dependent and independent environmental factors. In this study, we used the official 1998–2003 inventory data on ungulate numbers from 462 forest districts and 23 national parks across Poland to determine the roles of various environmental factors in shaping country-wide spatial patterns of ungulate abundances. Spatially explicit generalized additive mixed models showed that different sets of environmental variables explained 39 to 50 % of the variation in red deer Cervus elaphus, wild boar Sus scrofa, and roe deer Capreolus capreolus abundances. For all of the studied species, low forest cover and the mean January temperature were the most important factors limiting their numbers. Woodland cover above 40–50 % held the highest densities for these species. Wild boar and roe deer were more numerous in deciduous or mixed woodlands within a matrix of arable land. Furthermore, we found significant positive effects of marshes and water bodies on wild boar abundances. A juxtaposition of obtained results with ongoing environmental changes (global warming, increase in forest cover) may indicate future growth in ungulate distributions and numbers.     

Trophic niche overlap and wild ungulate consumption by red fox and wolf in a mountain area in Italy

Red fox (Vulpes vulpes) and wolf (Canis lupus) are two widespread opportunistic predators living in simpatry in many areas. Nonetheless, scarce information are available on their trophic interactions. We investigated food habits of these two carnivores in a mountain area in Italy and assessed the extent of their trophic niche overlap, focusing on the consumption of wild ungulates. Thereby we analyzed the content of 669 red fox scats and 253 wolf scats collected between May 2008 and April 2009. Red foxes resulted to have a more than three times higher niche breadth than wolves. Vegetables, small mammals, wild ungulates, and invertebrates were major items (altogether 92% of volume) of the red fox annual diet. On the contrary wolf annual diet relied on wild ungulates (94% of volume) with wild boar (Sus scrofa) being the main food item. The degree of trophic niche overlap between the two species was found to be low (Pianka's O = 0.356). Diet variation between the warm and the cold seasons was limited in both species, and higher in red fox than in wolf. The two canids appeared to use wild ungulates unevenly being the former more selective for younger preys, smaller in size (newborn piglets and roe deer Capreolus capreolus fawns), whereas the latter exhibited a preference for medium-sized and large ungulates (10–35 kg wild boar and adult roe deer). Even if wild ungulates represent the main shared food category, the different use of age/weight classes by the two predators, together with their possible consumption as carrions by red fox, suggests a very limited trophic competition between wolf and red fox.This study represents a contribution to the knowledge of trophic interaction in predator–prey systems where sympatric carnivores are present.     

Comparing patterns of human harvest and predation by Eurasian lynx Lynx lynx on European roe deer Capreolus capreolus in a temperate forest

Human harvest is the most important mortality factor for wild ungulates in Europe and can affect several aspects of ungulate biology. There is a growing concern about possible negative side effects of human harvest. To better understand the differences between human and natural mortality, we compared the extent, age and sex structure, nutritional condition, spatial and temporal distribution of human harvest, and natural predation by the Eurasian lynx Lynx lynx on the European roe deer Capreolus capreolus, the most abundant wild ungulate in Europe. Compared to the human harvest, lynx were less likely to kill fawns and yearlings than adults, and among adult deer, lynx were more likely to kill females. The proportion of roe deer with fat-depleted bone marrow was higher among lynx prey than among harvested animals. Average lynx kill rate was estimated to 47.8 roe deer per year, and lynx predation was considerably lower than the human harvest in the same area. While human harvest increased with higher roe deer density, lynx predation was similar across the gradient of roe deer densities. Comparison with other countries indicated that differences between human harvest and natural mortality of ungulates vary considerably in different parts of Europe. Variation in hunting practices and, even more importantly, carnivore predation may have an important role in buffering unwanted side effects of harvest of wild ungulates.     

Using basic plant traits to predict ungulate seed dispersal potential

Habitat fragmentation contributes to the decline of plant species by decreasing gene flow among populations. Restoring connectivity among habitat patches is therefore a major issue for plant conservation. However, deciding where to focus restoration efforts requires identifying suitable dispersers for each target plant species. We collected data from the literature on wild and domesticated ungulates, known to be effective seed dispersers, and on the plants they dispersed in Europe via epi‐ and/or endozoochory. We performed a systematic literature review to identify plant and animal traits relevant for seed dispersal. We first modeled the relationships between epi‐ or endozoochory and a priori selected plant traits (diaspore releasing height, length, shape and morphology, and habitat openness). The differences we underlined between the two dispersal mechanisms justified splitting our analyses accordingly. Then, for each dispersal mechanism, we asked whether basic plant traits could be used to predict specific traits of ungulates as endozoochorous or epizoochorous seed dispersers. We modeled the relationships between a priori selected ungulate traits for epizoochory (habitat openness, shoulder height, hair curliness, and hair length) and for endozoochory (habitat openness, body mass, feeding type and digestive system) and plant traits. Plant habitat openness and diaspore morphology were the predictors that most often explained differences among ungulates for epizoochory, whereas plant habitat openness and diaspore releasing height most often explained differences for endozoochory. Our trait‐based predictive models can help improve our ability to propose more precise management decisions for the conservation of plant populations worldwide by taking into account ungulate dispersers.     

Population dynamics in a guild of four Mediterranean ungulates: density‐dependence,environmental effects and inter‐specific interactions

Population fluctuations in ungulates are driven by both intrinsic and extrinsic factors. Available information, however, mainly refers to arctic, temperate and African ungulate populations, while the dynamics of Mediterranean species, exposed to a milder climate, is known to a much lesser extent. Here we studied the population dynamics of four wild ungulate species in the Castelporziano Preserve near Rome, Italy, as obtained from detailed bag counts from hunting drives during the period 1878–1986: the Italian roe deer Capreolus capreolus italicus, the Maremma wild boar Sus scrofa majori (both endemic to Italy), the native red deer Cervus elaphus, and the alien fallow deer Dama dama. We also considered the effects of the presence of another alien ungulate, the nilgai Boselaphus tragocamelus. This ungulate community experienced an accidental ‘removal experiment’ when, during World War II, red deer and nilgai were exterminated. This event and the length of the time series allowed us to test two main hypotheses: 1) that the complexity level of the ungulate community affects the strength of intra‐ and inter‐specific competition; and 2) that in Mediterranean environments intra‐ and inter‐specific interactions are stronger than climate forcing. Statistical methods ranged from state‐space‐modelling, GLM analysis and structural equation models. The results indicated that direct intra‐specific density dependence played a relevant role for all species, and was stronger after the removal. A complex pattern of species interactions was however revealed; fallow deer had a negative effect on roe deer population, while roe deer had an apparent positive effect on red deer and wild boar, possibly mediated by environmental factors. Nilgai appeared to facilitate all deer species. The results of the analysis also confirmed that at present climate appears to play a minor role with respect to density dependence; however, the increasing aridity of the Mediterranean area could change this picture in coming decades.     

Seed dispersal by ungulates as an ecological filter: a trait‐based meta‐analysis

Plant communities are often dispersal‐limited and zoochory can be an efficient mechanism for plants to colonize new patches of potentially suitable habitat. We predicted that seed dispersal by ungulates acts as an ecological filter – which differentially affects individuals according to their characteristics and shapes species assemblages – and that the filter varies according to the dispersal mechanism (endozoochory, fur‐epizoochory and hoof‐epizoochory). We conducted two‐step individual participant data meta‐analyses of 52 studies on plant dispersal by ungulates in fragmented landscapes, comparing eight plant traits and two habitat indicators between dispersed and non‐dispersed plants. We found that ungulates dispersed at least 44% of the available plant species. Moreover, some plant traits and habitat indicators increased the likelihood for plant of being dispersed. Persistent or nitrophilous plant species from open habitats or bearing dry or elongated diaspores were more likely to be dispersed by ungulates, whatever the dispersal mechanism. In addition, endozoochory was more likely for diaspores bearing elongated appendages whereas epizoochory was more likely for diaspores released relatively high in vegetation. Hoof‐epizoochory was more likely for light diaspores without hooked appendages. Fur‐epizoochory was more likely for diaspores with appendages, particularly elongated or hooked ones. We thus observed a gradient of filtering effect among the three dispersal mechanisms. Endozoochory had an effect of rather weak intensity (impacting six plant characteristics with variations between ungulate‐dispersed and non‐dispersed plant species mostly below 25%), whereas hoof‐epizoochory had a stronger effect (eight characteristics included five ones with above 75% variation), and fur‐epizoochory an even stronger one (nine characteristics included six ones with above 75% variation). Our results demonstrate that seed dispersal by ungulates is an ecological filter whose intensity varies according to the dispersal mechanism considered. Ungulates can thus play a key role in plant community dynamics and have implications for plant spatial distribution patterns at multiple scales. Synthesis Plant communities are often dispersal‐limited and zoochory can be an efficient mechanism for plants to colonize new patches of potentially suitable habitat. Our analysis is the first synthesis of ungulate seed dispersal that compares characteristics from both non‐dispersed and dispersed diaspores, distinguishing the three zoochory mechanisms ungulates are involved in: endozoochory, hoof‐epizoochory and fur‐epizoochory. We confirmed that seed dispersal by ungulates is an ecological filter whose intensity increases from endozoochory, then hoof‐epizoochory to finally fur‐epizoochory. By filtering seed traits through dispersal, ungulates can thus play a key role in plant community dynamics and have implications for plant spatial distribution patterns at multiple scales.     

Rewilding traditional grazing areas affects scavenger assemblages and carcass consumption patterns

The abandonment of traditional livestock farming systems in Mediterranean countries is triggering a large-scale habitat transformation, which, in general, consists of the replacement of open grazing areas by woodlands through non-managed regeneration. As a consequence, wild ungulates are occupying rapidly the empty niche left by domestic ungulates. Both types of ungulates represent the main trophic resource for large vertebrate scavengers. However, a comparison of how vertebrate scavengers consume ungulate carcasses in different habitats with different ungulate species composition is lacking. This knowledge is essential to forecast the possible consequences of the current farmland abandonment on scavenger species. Here, we compared the scavenging patterns of 24 wild and 24 domestic ungulate carcasses in a mountainous region of southern Spain monitored through camera trapping. Our results show that carcasses of domestic ungulates, which concentrate in large numbers in open pasturelands, were detected and consumed earlier than those of wild ungulate carcasses, which frequently occur in much lower densities at more heterogenous habitats such as shrublands and forest. Richness and abundance of scavengers were also higher at domestic ungulate carcasses in open habitats. Vultures, mainly griffons (Gyps fulvus), consumed most of the carcasses, although mammalian facultative scavengers, mainly wild boar (Sus scrofa) and red fox (Vulpes vulpes), also contributed importantly to the consumption of wild ungulate carcasses in areas with higher vegetation cover. Our findings evidence that the abandonment of traditional grazing may entail consequences for the scavenger community, which should be considered by ecologists and wildlife managers.     

Ungulate browsers promote herbaceous layer diversity in logged temperate forests

Ungulates are leading drivers of plant communities worldwide, with impacts linked to animal density, disturbance and vegetation structure, and site productivity. Many ecosystems have more than one ungulate species; however, few studies have specifically examined the combined effects of two or more species on plant communities. We examined the extent to which two ungulate browsers (moose [Alces americanus]) and white‐tailed deer [Odocoileus virginianus]) have additive (compounding) or compensatory (opposing) effects on herbaceous layer composition and diversity, 5–6 years after timber harvest in Massachusetts, USA. We established three combinations of ungulates using two types of fenced exclosures – none (full exclosure), deer (partial exclosure), and deer + moose (control) in six replicated blocks. Species composition diverged among browser treatments, and changes were generally additive. Plant assemblages characteristic of closed canopy forests were less abundant and assemblages characteristic of open/disturbed habitats were more abundant in deer + moose plots compared with ungulate excluded areas. Browsing by deer + moose resulted in greater herbaceous species richness at the plot scale (169 m²) and greater woody species richness at the subplot scale (1 m²) than ungulate exclusion and deer alone. Browsing by deer + moose resulted in strong changes to the composition, structure, and diversity of forest herbaceous layers, relative to areas free of ungulates and areas browed by white‐tailed deer alone. Our results provide evidence that moderate browsing in forest openings can promote both herbaceous and woody plant diversity. These results are consistent with the classic grazing‐species richness curve, but have rarely been documented in forests.     

Differential long-term impacts of a management control program of axis deer and wild boar in a protected area of north-eastern Argentina

Exotic ungulates are among the top global invasive mammals and a threat to biodiversity. Axis deer (Axis axis) and wild boar (Sus scrofa) are of increasing concern in multiple regions. A management program reduced wild boar abundance and soil damage below target levels through controlled still shooting from watchtowers and dog-hunting performed by recreational hunters at El Palmar National Park, Argentina. Here we assess program impacts on axis deer over a 10-year period in which 2380 deer were dispatched, and document two largely unexpected outcomes: increasing axis deer abundance toward a plateau, and a strong inverse correlation between deer and wild boar numbers. Unlike the initial steep decline and subsequent stabilization of wild boar, deer abundance indexed by standardized catch-per-unit-effort increased at 37.6% per year over 0–5 years post-intervention (YPI) and stabilized from 7 YPI on when still-shooting effort averaged 948 hunting party-hours per quarter. Deer catch was non-linearly related to still-shooting effort. Timing of deer and boar catches did not differ significantly regardless of sex, season and YPI. Catch-per-unit-effort indices and nightly spotlight deer counts showed similarly increasing trends. The fraction of older adult deer declined over 0–4 YPI and remained stable thereafter. Sex ratios were consistently skewed toward males only among older adults. Failure to reduce deer abundance may be explained by several major processes: protracted exponential growth of the deer population after park invasion; deer regional expansion with increasing immigration; insufficient sex- and stage-biased hunting mortality, and competitor (and perhaps predator) release from wild boar.     

Habitat niche relationships within an assemblage of ungulates in Bardia National Park,Nepal

Species co-exist in their individual niches by partitioning resources. The main prey of the big predators – wild boar, chital, and sambar are sympatric in tropical south-Asia. In order to obtain a better understanding of their habitat use and habitat niche relationships, we used an indirect sign (faecal pellets and rooting by wild boar) survey along line transects in Bardia National Park. Ground cover, in particular grass cover for wild boar and chital, and shrub cover for chital and sambar positively affected the occurrence of ungulate signs. We found a niche differentiation between wild boar, which preferred hill sal forest and grassland, and chital and sambar, which avoided hill sal forest and grassland. Habitat niche overlap between chital and sambar in open habitat was clearly evident during the dry season. It is suggested that the fine grained habitat mosaics in space are the underlying causes of the coexistence of the ungulates studied. Therefore, focus on maintaining the habitat mosaic throughout the regions of the species’ distribution may prove successful for effective conservation management.